NOTES ON MEXICAN ACTINOTE (NYMPHALIDAE: ACRAEINAE) AND THEIR RELATIVES, WITH DESCRIPTION OF A NEW SUBSPECIES

Journal of the Lepidopterists' Society 32(4), 1978, 261-272 NOTES ON MEXICAN ACTINOTE (NYMPHALIDAE: ACRAEINAE) AND THEIR RELATIVES, WITH DESCRIPTION OF A NEW SUBSPECIES JACQUELINE Y. MILLER AND LEE D. MILLER Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580 ABSTRACT. The application of names described for South American species to their Central American, and especially Mexican, counterparts has led to great confusion in the literature. Actinote stratonice oaxaca is described from Oaxaca, Mexico; this insect had been reported previously as the nominate subspecies. Three species of the Actinote thalia group, A. calderoni, lapitha and thalia are illustrated, redescribed and discussed. A key for the separation of the three species is provided. The first two species are recorded from Mexico-calderoni had been previously misidentified as lapitha from there. One name, A. lapitha zilchi Franz and Schroder, is synonymized to calderoni. Collecting in Mexico over the past forty years has yielded many butterfly species that were previously unknown from there, not a few of which were totally unexpected. These unexpected taxa have created many problems, usually for one of two reasons: 1) the butterfly was an already described Central or South American species and was described as new from Mexico because of a lack of comparative material or an ignorance of the pertinent literature; or 2) many species (especially those figured and described in Seitz) incorrectly have been ascribed to the Mexican fauna, again because of a lack of adequate comparative material. Both situations are well demonstrated in the Nymphalidae: Acraeinae. The Hoffmann (1940) catalog lists only four species of this subfamily within the borders of Mexico, but recent collecting has uncovered one that has been misidentified in collections and in correspondence-the one with which the first-mentioned species had been confused and an undescribed subspecies of a well-known South American insect. In the hope of unravelling the confusion in this small subfamily (within the Mexican borders), we offer these notes. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies Figs. 1-6 Male. Head, thorax and appendages black; abdomen black with a reddish-brown midsternal line. Upper surface of wings similar to that of the nominate subspecies, but paler; and with the following differences: forewing totally black anteriad of cell (partially reddened in other subspecies); dark marking at end of forewing cell much smaller than in other subspecies and black area at base of forewing cell and along inner margin more restricted than in other populations. Under surface pattern also paler than in s. stratonice with forewing differences as noted for upper surface

262 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Figs. 1-4. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies. 1-2, Holotype ~, upper (1) and under (2) surfaces; MEXICO: OAXACA: Candelaria Loxicha, 550 m. 3-4, Paratype <;', upper (3) and under (4) surfaces; same locality. Both specimens in Allyn Museum of Entomology. and in addition the pale orange patches in the forewing cell area darkened basad, blending to tawny-yellow distad; black areas admixed with pale yellow scales, especially in base hindwing. ~ genitalia as illustrated, differing from those of other stratonice in only minor respects. Length of forewing of Holotype ~ 24.4 mm, those of eleven of the ~ Paratypes range from 26 to 31 mm. Female. Differs from the <;' of nominate stratonice in the same manner as does the ~, but additionally the basal black areas of the forewing encompass the proximal third of the cell and below it along the inner margin, and all of the black areas below are completely suffused with yellow scaling. One <;' Paratype has a hindwing supernumerary vein off Rs on the right side. <;' genitalia as illustrated and comparing well with those of other subspecies. Lengths of forewings of 12 <;' Paratypes range from 28.2 to 38 mm. Specimens examined. Described from 25 specimens, 13 males and 12 females, from the state of Oaxaca, Mexico. Types. HOLOTYPE ~: MEXICO: OAXACA: Candelaria Loxicha, 550 m, 8.ix. 1969 (E. C. Welling). PARATYPES: all MEXICO: OAXACA: same locality as HoIotype, 1 ~ 15.xi.1967, 1 ~ 15.ix.1968, 1 <;' 21.vii.l970, 1 ~ 1 <;' 27.viii.l970, 1 <;' 21.vii.l973 (all E. C. Welling M.); El Portillo del Rayo, Candelaria Loxicha, 1 ~ 3<;' 17.xi.l967.(all E. C. Welling M.), 6~ 4<;' 18.vii.l976 (ali de Ia Maza family); Rio Molina, Mpio. Suchistepec, 2200 m., 1 <;' 1O.x.1967 (E. C. Welling M.); San Jose Pacifico, Mpio. Rio Hondo, 2400 m., 1 ~ 9.x.1967 (E. C. Welling M. ); Puente

1 VOLUME 32, NUMBER 4 263 Figs. 5-6. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies. 5, $ genitalia of Paratype; MEXICO: OAXACA: Candelaria Loxicha; preparation M-3630 (Jacqueline Y. Miller). 6, 'i? genitalia of Paratype; same locality; preparation M-3602 (Jacqueline Y. Miller). del Guajolote, J alatengi, 1 'i? 21.iii.1975 (E. Fernandez), 1 $ 21.xi.1975 (de la Maza family); Dos de Mayo, 1$ 18.v.1976 (de la Maza family). Disposition of type series: Holotype $, two (1) and four 'i? Paratypes in the collection of the Allyn Museum of Entomology; two $ and three 'i? Paratypes in the collection of E. C. Welling M. and eight (1) and five 'i? Paratypes in the de la Maza collection. Remarks. The sub specific name refers to the state from whence the new subspecies came. A. stratonice is recorded in Seitz from the Sierra Madre de Santa Marta of Colombia and southward through the mountains of Venezuela and into Ecuador. The present subspecies was first recorded in the literature from Mexico by de la Maza R. and de la Maza E. (1975), but at the time they did not recognize it as a separate subspecies. The subspecies oaxaca is characterized by its overall dull coloration and by the suffusion of the under surface with pale yellow scales. In none of the other subspecies of stratonice do these characters appear. The disjunct distribution of stratonice, with a Mexican subspecies widely separated from its nearest relatives, is indeed intriguing.

264 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Figs. 7-10. Actinote calderoni Schaus. 7-8, 3, upper (7) and under (8) surfaces; MEXICO: CHIAPAS: Mapastepec. 9-10, 9, upper (9) and under (10) surfaces; same locality. Both specimens in Allyn Museum of Entomology. The Identity of the Mexican Actinote thalia Group Species Pale Mexican specimens of thalia group Actinute in most collections usually have been identified as A. lapitha (Staudinger). Comparison of most Mexican material with Staudinger's description and subsequent literature citations suggest that these specimens could not be true lapitha. Accordingly, we searched museum and private collections and the literature for documented material that might shed light on the correct name for the Mexican insects. The results were surprising, and to avoid future confusion the following redescriptions are offered to aid in the identification of these butterflies. Actinote calderoni Schaus, 1920 Figs. 7-12 Actinote calderoni Schaus, 1920: 434 (TL-Anteos, EI Salvador). = Actinote lapitha zilchi Franz and Schroder, 1954: 80; fig. 2 (TL-Km. 30, Sonsonate Rd., La Libertad, EI Salvador). [New Synonymy.] Male. Head, thorax and first two abdominal segments dark brown covered with a few fuscous and tawny dorsal hairs; remaining abdominal segments naked

VOLUME 32, NUMBER 4 265 @ Figs. 11-12. Actinote calderoni Schaus. 11, t gcnitalia; MEXICO: CHIAPAS: Mapastepcc; preparation M-3633 (Jacqueline Y. Miller). 12, <j! sterigma; same locality; preparation M-3601 (Jacqueline Y. Miller). dorsad; thorax and first two abdominal segments covered with a few tawny ventral scales; last abdominal segments ventrally naked; pleural line buff. Palpi tawny with a few black scales along inner surface only. Antennae and legs black. Upper surface of wings very thinly scaled with white, giving a dusky, off-white appearance; forewing smoky at apex and marginally to CU2-2A; veins slightly darkened and interneural spaces with single smoky stripes. Under surface of wings similar, but a faint, dark comma-shaped marking lies from M,,-Cut to CUt-CU2 of the forewing and numerous long, dark hairs lie on the veins, especially of the hindwing. t genitalia as illustrated. Forewing lengths of the t examples examined range from 21.5 to 24.5 mm. Female. Similar to $, but paler, especially the forewing smoky markings above and below. <j! genitalia as illustrated, generally characterized by having the sterigma more sculptured than in related species. Lengths of forewings of <j! specimens at hand range from 25 to 30.5 mm. Types and specimens examined. We have examined 15 specimens, seven males and eight females, from the following localities: EL SALVADOR: Anteos, 1 t (HOLOTYPE, USNM), It (possible PARATYPE, CM). GUATEMALA: Tiquisate, 28.vi.1947, 1 <j! (AMNH). MEXICO: CHIAPAS: Mapastepec, various dates, 1939-1959, 2t 5<jl (AME), 3$ H (AMNH); San Jeronimo, 600 m., 17.vii.1975, H (E. C. Welling M.). The records from Mexico and Guatemala are apparently the first for either country. The present insect has been masquerading in Mexican collections for years as A. lapitha (Staudinger), a species that is abundantly distinct. Not only have workers on Mexican butterflies been confused on the identity of calderoni, but also those in other parts of Central America. The fact that Schaus' (1920) description appeared in an entomologically obscure journal has resulted in the paper never being cited previously by workers on Actinote. Were the "Fifty-Year Rule" still in effect in

266 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Figs. 13-16. Actinote lapitha (Staudinger). 13-14, is, upper (13) and under ( 14) surfaces; MEXICO: CHIAPAS: Tierra Blanca, Mpio. La Trinitaria, 1500 m. (E. C. Welling M. collection). 15-16, 'i', upper (15) and under (16) surfaces; no locality data (AMNH collection). the International Code of Zoological Nomenclature, the name calderoni could be ignored with impunity, but happily this rule was repealed a few years ago, so we must return to the oldest name. Franz and Schroder (1954) had obviously not seen the Schaus description, but their excellent figure of the type of A. lapitha zilchi is referable to specimens of calderoni that we have seen (including the Holotype), and since the two taxa were described from within 100 km. of one another, it is evident that they n~present the same species. Actinote lapitha (Staudinger), 1888 Figs. 13-18 Acraea lapitha Staudinger, 1888: 82 (TL-"Chiriqui"). Male. Head, thorax and abdomen clothed with black dorsal hairs; head, palpi and most of thorax also clothed with black ventral hairs; small patch of tawny scales on meso- and meta thoracic preepisterna and an additional such patch on metathoracic epimeron; abdomen clothed with admixed fuscous, tawny and buff scales; pleural line tawny and buff only. Antennae and legs black. Ground color of forewing above translucent and tawny with margins outlined in dull gray-brown, especially at apex; prominent gray-brown transverse marking from end cell to Cu,-2A, interspersed with

VOLUME 32, NUMBER 4 267 1e:===3J~~.::~...:... ~?" Figs. 17-18. Actirwte lapitha (Staudinger). 17, ~ genitalia; MEXICO: CHI APAS: Tierra Blanca, Mpio. La Trinitaria; preparation M-3416 (Jacqueline Y. Miller). 18, ';> sterigma; no data; preparation M-1657 (Jacqueline Y. Miller). tawny scaling from end cell to M,-Cul; veins and interneural striping prominent, gray-brown. Hindwing above also translucent tawny, dull gray-brown at costa and along margin; veins heavily darkened with gray-brown and interneural markings of same color short and heavy. Forewing below similar to upper surface, but margins and apex not so dark and with an additional dull gray-brown bar across middle of cell. Hindwing below as above with a faint gray-brown cell end marking from costa to base of M, and more prominent veinal and interneural blackish-brown striping. Fringes of both wings blackish on both surfaces. ~ genitalia as illustrated. Lengths of forewings of the three ~ specimens examined range from 22.8 to 23.6 mm. Female: Similar to ~, but thorax sparsely clothed with tawny scales, markings of all wings paler, but extra-discal band of forewing more prominent and base of cell of same wing somewhat overscaled with fulvous. ';> genitalia as illustrated; sterigma somewhat heavier than that of the next species and not quite so ornamented as in calderoni. Lengths of forewings of the two ';> examples before us 20.2 and 23.1 mm. Types and specimens examined. We have seen two females and three males of this insect. PANAMA: Jicaron Island, 14-15.i.1902, 1 ~ (BMNH). COSTA RICA: Puerto Golfito, 4.vii.l965, 1 ~ (Gordon B. Small, Jr. collection). MEXICO: CHI APAS: Tierra Blanca, Mpio. La Trinitaria, 1500 m., 15.ix.1972, 1 ~ (E. C. Welling M.) No Data, 2';> (AMNH). Evidently the Costa Rican record is a new, but not unexpected one. Hoffmann (1940: 672) lists lapitha from "Tierra caliente de la costa del Pacifico de Chiapas," no true specimens of that species are in the Hoffmann collection in tbe AMNH. All of the specimens in Hoffmann's material were calderoni, and one of these bore a determination label in Hoffmann's hand of "Actinote lapitha Staudinger." Since Hoffmann obviously confused lapitha with calderoni, we feel that Mr. Welling's specimen of the former is the first authentic record from Mexico. A. lapitha was described from the Chiriqui region of Panama, but the type specimen is apparently no longer extant, perhaps having been

268 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Figs. 19-22. Actinote stlbhyalina (Staudinger). 19-20, ~, upper (19) and under (20) surfaces; PERU: Rio Cachiyacu, Iquitos (BMNH). 21-22, Lectotype «(see designation in text), upper (21) and under (22) surfaces; PERU: Yurimaguas (ZMHU). destroyed during World War II (H. J. Hannemann, pets. comm.). We have been unable to locate an authentic Chiriqui specimen in more recent collections; hence, we do not designate a Lectotype here. Actinote subhyalina (Staudinger), 1888 Figs. 19-24 Acraea subhyalina Staudinger, 1888: 81; pi. 32 (TL-Yurimaguas, Peru). Description. Sexes similar: Head, thorax and abdomen above and below blackish-brown clothed with fuscous to buff scales. Palpi blackish-brown; legs brown and antennae dull reddish-brown. Forewing above buff, translucent and laved with fulvous toward base (darkest in ~); apex and margin dull brown; veins darkened; transverse markings: a jagged blackish-brown one across cell about %, distance from base to end and a second one from costa across end cell to anal margin (not prominent below CUt in ~). Hindwing above buff, translucent, with darker veins; margin outlined in dull brown; interneural striping in anal region; prominent dark brown bar from costa to CUt-CU, and a dull brown mark across distal end of cell. Under surface of wings as above, but dull brown markings slightly overscaled with buff, hindwing distal band more diffuse and an additional dull brown stripe in hindwing cell Sc+Rt-Rs. Lengths of forewings of all specimens examined ranged from 20 to 22 mm. Genitalia of ~ and «as illustrated. Specimens examined. We have been able to examine only a single ~ and six «specimens. PERU: Rio Cachiyacu, Iquitos, [18193, Stuart, 1~ 3«(BMNH); Yurimaguas, 1«(ZMHU); No data, 2«(CM).

VOLUME 32, NUMBER 4 269 @ jjj;;--:~a±... ~...@?~ Figs. 23-24. Actinote subhyalina (Staudinger). 23, is genitalia; PERU: Rio Cachiyacu, Iquitos; preparation M-3403 (Jacqueline Y. Miller). 24, 'i' genitalia of Lectotype; PERU: Yurimaguas; preparation M-3411 (Jacqueline Y. Miller). Staudinger (1886: 81) described A. subhyalina from 12 'i' specimens taken at Yurimaguas. Certainly the figured 'i' from Berlin was one of Staudinger's syntypes, and the two specimens from CM may have been, but in the case of the latter two specimens this cannot be ascertained with precision. One of the CM specimens bears the label "Acraea subhyalina/ from Dr. O. Staudinger/1885," a date that was three years before the description; the second specimen bears a number (Staudinger's?) only, "386." The specimen received from the ZMHU is definitely from the type locality, was in the Staudinger collection and has been labelled as "Origen" by Staudinger or someone subsequent to him. It is the logical candidate for designation as the Lectotype of the name, and we have so labelled it, affixing a red, partially printed, partially handwritten (italics) label to it: "Lectotype/ Acraea/ subhyalina/ Staudinger, 1888/ designated by Jacqueline Y. Miller/ & Lee D. Miller, 1977." DISCUSSION The impetus for this project was a series of seven specimens in the Allyn Museum collection from Mapastepec, Chiapas, Mexico. Examination of these specimens revealed that while they were closely related to A. lapitha (the name associated with them), they were abundantly distinct. Letters for additional specimens brought two from the AMNH that were in agreement with the original description of lapitha (but without data) and five more of the odd one, four of which were from the

270 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Hoffmann collection, taken by Dr. Escalante. A later trip to CM yielded one more specimen and the first clue to the identity of the Mexican material that we had. The additional label on the CM specimen stated that it was "Actinote calderoni Schaus," but at the time we were unable to find the original description or any reference to it, and we still assumed that the name might have been a manuscript name only. Rechecking the series and the type collection at the USNM yielded not only the type specimen of calderoni, but also finally the reference to the original description of this elusive name. The confusion did not end there, though. The two true lapitha from the AMNH had a fulvous basal flush on the forewing, thus resembling subhyalina. This led us to wonder if the specimen figured by Staudinger ( 1888) was subhyalina or lapitha. Letters to the BMNH and the ZMHU brought additional specimens of Staudinger's insects and some very helpful information. All three species are rare in collections, especially the males. Since we encountered such difficulty in making determinations in the thalia group, we present the following key to aid other workers to the species treated here. 1. Forewing above with dark transverse band at end cell from costa to near anal angle 2. I'. Forewing with no such band; Mexico to El Salvador calderoni Schaus. 2. Hindwing above with prominent dark discal markings at end of cell; Peru subhyalina (Staudinger). 2'. Hindwing above without prominent marking at end cell; Mexico to Panama lapitha (Staudinger). Essentially the key characters for the separation of the species in this complex are the dark bars across the cells of both wings. A. subhyalina shows these bars on the upper surfaces of both wings, A. lapitha has only the one on the forewing and A. calderoni has neither. Seemingly the orange flush at the base of the upper forewing should be diagnostic, but whereas it is prominent in most subhyalina, it also appears in some female lapitha, hence it is diagnostic of neither. The male genitalia (Figs. 11, 17, 23) are similar, but subtly different, in all three species. The valvae are elongated and slightly curved dorsad in calderoni, whereas they are squarecut posteriad in both lapitha and subhyalina. The saccus of subhyalina is much more elongate than is that of either of the other two species. Most of the differences between females of these species lie in the sterigmal region. The posterior margin of the lamella postvaginalis is U-shaped and narrow in lapitha, U-shaped and expanded in subhyalina and even more enlarged and W-shaped in calderoni. The entire opening

VOLUME 32, NUMBER 4 271 of the ostium bursae is darkly sclerotized in a narrow ring in lapitha, a somewhat broader darkly sclerotized ring in subhyalina but only darkly sclerotized in four separate areas around the opening in calderoni, but the area around the ostium itself is lightly sclerotized. The genital capsule is larger in calderoni than in the other species. The tarsi, as is true of all members of the thalia group, are asymmetrical and comparable one with another. Geographically A. subhyalina can be immediately separated from lapitha and calderoni, none of the three species being found in Colombia or Ecuador, as far as we know. It would not be surprising to see subhyalina from at least the latter country, and there might be sympatry between subhyalina and lapitha in Colombia. A. lapitha, as recorded here, has a much more extensive range than previously believed, and calderoni is not restricted to EI Salvador. Either of these species, or both, may well be found in Honduras and Nicaragua. We hope that this paper will encourage others to try to fill in the distributional blanks for this interesting group. ACKNOWLEDGMENTS Weare deeply indebted to the following for loan of material and for access to their collections (abbreviations) of Actinote: Dr. F. H. Rindge, American Museum of Natural History (AMNH); Mr. H. K. Clench, Carnegie Museum of Natural History (CM); Mr. W. D. Field, National Museum of Natural History (USNM); Mr. P. R. Ackery, British Museum (N atural History) (BMNH); Dr. H. J. Hannemann, Zoologische Museum del' Humboldt U niversitiit (ZMHU); Dr. T. Escalante and Messrs. R. de la Maza R. and J. de la Maza E., Mexico, D. F., Mexico; Mr. E. C. Welling M., Merida, Yucatan, Mexico and Mr. G. B. Small, Jr., Balboa, Canal Zone. The Allyn Museum of Entomology is abbreviated AME in part of the paper. Mr. Field and Dr. J. F. G. Clarke of the USNM were instrumental in obtaining a copy of the Schaus paper for us. We also thank Mr. S. R. Steinhauser for pointing out the Franz and Schroder paper to us and to Mr. H. W. Dybas of the Field Museum for obtaining a copy of it for our use. Special thanks are due Mr. A. C. Allyn of this institution for taking the photographs contained herein and for reading the manuscript. LITERATURE CITED FRANZ, E. & H. SCHRODER. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergische BioI. 35: 75-87; figs.

272 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY HOFFMANN, C. C. 1940. Catalogo sistematico y zoogeografica de los Lepidopteros mexicanos. 1a. Parte. Papilionoidea. An. lnst. BioI. 11: 639-739. JORDAN, K. 1924. Acraeidae. In Seitz, A., Macrolepidoptera of the World 5: 358-374. DE LA MAZA R., R. & J. DE LA MAZA E. 1975. Adiciones al Catalogo de Lepidopteros mexicanos. Rev. Soc. Mexicana Lepid. 1: 62-63; figs. SCHAUS, W. 1920. Descriptions of two new species of butterflies from tropical America. J. Washington Acad. Sci. 10: 434-435. STAUDINGER, O. 1888. In Staudinger, O. and E. Schatz, Exot. Schmett. (1): 81-82; PI. 32.